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Seed Morphology and Anatomy of Hypericum Majus (A biodiv. res. Conserv. 55: 7-14, 2019 BRC www.brc.amu.edu.pl Doi 10.2478/biorc-2019-0013 Submitted 19.07.2019, Accepted 30.09.2019 Seed morphology and anatomy of Hypericum majus (A. gray) britton Piotr Szkudlarz1*, Zbigniew Celka1, Stanisław rosadziński# & Maria Katarzyna Wojciechowicz2 1Department of Plant Taxonomy, Faculty of biology, Adam Mickiewicz University, Poznań, Uniwersytetu Poznańskiego 6, 61-614 Poznań, Poland; orCiD: PSz – http://orcid.org/0000-0002-4915-2721, ZC – http://orcid.org/0000-0002-5344-8939 2Department of general botany, Faculty of biology, Adam Mickiewicz University in Poznań, Uniwersytetu Poznańskiego 6, 61-614 Poznań, Poland; orCiD: MKW – http://orcid.org/0000-0001-9602-2535 *corresponding author (e-mail: [email protected]) Abstract. Hypericum majus (A. gray) britton is an alien species for europe. it originates from north America and, in recent years, has been spreading in europe. Presently, this species is found in germany, France, italy, bosnia and herzegovina, and belarus. in Poland, it was noted for the first time in 2017 in western part of the country. Thus far, no detailed morphology and anatomy of H. majus seeds based on light and electron microscopy studies has been presented. Considering the fact that seeds provide important taxonomical information and the discussed species has not been investigated in this respect, in this research, an attempt was made to broaden the present knowledge. investigations were conducted based on samples originating from three localities in Central europe (Poland). The obtained results showed that H. majus seeds are very small – about 0.5 mm in length. in the SeM image, their surface is reticulate, typical of majority of species in this genus. Anatomical analysis showed that seed coat of matured seeds is very delicate, composed of two cell layers with strongly lignified walls. hylogeny Key words: carpology, greater Canada St. John's-wort, hypericaceae, Poland, seed sculpture, seed coat P D n A 1. Introduction rianum Wight & Arn. were found in crops (Mirek et al. 2002; Jackowiak et al. 2013, 2017). y The world flora of the genus Hypericum l. com- In the flora of Poland, a new, non-native species M prises, according to different sources, from about 370 of Hypericum, which spreads also in many european to 520 species (Stevens 2007; Mabberley 2008; nürk countries, is H. majus (A. gray) britton (nobis et al. & blattner 2010; Crockett & robson 2011; The Plant 2018). This taxon originates from north America. it list 2013; hypericum online). in europe, this genus is is a perennial plant that grows up to 0.7 m in height xono represented, depending on taxonomical approach, by and develops one or several quadrangular stems. leaf 56-61 species (Mabberley 2008; robson 2005), with blades are lanceolate or elliptic (longitudinal) with entire ta 13 of these taxa observed in Poland to date (Mirek margin. inflorescence is apical and may consist of even et al. 2002). The majority of species occurring in Po- 30 flowers. Flowers are stellate, 6-7 mm in diameter, y, land are native taxa (H. elegans Stephan ex Willd., petals golden-yellow (3.5-6.5 × 0.8-1.5 mm). Stamens T H. hirsutum l., H. humifusum l., H. maculatum Crantz, 12-21, ovary 1.5-2.0 × 0.8-1.0 mm, ovoid to ellipsoid; H. montanum l., H. perforatum l., H. pulchrum l., and styles 3; stigmas narrowly capitate (hypericum online). bili H. tetrapterum Fr.) or ephemerophytes (H. gymnanthum The floristic composition ofH. majus phytocoenoses engelm. & A. gray, H. mutilum l., and H. veronense matches phytosociological preferences of this species in A Schrank). Furthermore, H. calycinum l. and H. hooke- southern and western europe (bouchard 1954; nezadal ri A # co-author prepared this publication in cooperation with the Department of Plant Taxonomy V ©Adam Mickiewicz University in Poznań (Poland), Department of Plant Taxonomy. All rights reserved. 8 Piotr Szkudlarz et al. Seed morphology and anatomy of Hypericum majus (A. gray) britton 1984; Prince & Aniotsbehere 2012; Milanović 2018). in information about the anatomical structure of seeds in Poland, optimal habitats of H. majus comprise periodi- this genus has been also very limited (Szabes & Moro- cally exposed pond bottoms, water body margins and zowa 1991; Matzk et al. 2001; Szkudlarz 2014). periodically flooded former gravel pits.i t forms dense In Poland, all native Hypericum species were phytocoenoses that hinder the development of small investigated in detail in terms of their seed morphology terophytes from the Isoëto durieui-Juncetea bufonii (Szkudlarz 2014; Szkudlarz & Celka 2016) and in case class. Furthermore, locally, this species poses a threat of H. elegans also anatomically (Szkudlarz 2014). to phytocoenoses from the Hydrocotylo-Baldellion al- A new species – H. majus, which spreads in western liance (e.g., Pilularietum globuliferae). in overgrown, Poland, has not been studied yet in this regard in europe. unused ponds, H. majus was also found in the com- Thus, the aim of this work was an analysis and detailed munities representing the Molinio-Arrhenatheretea, description of morphological and anatomical structure Phragmitetea australis and Scheuchzerio-Caricetea of the seed coat in H. majus. fuscae classes (nobis et al. 2018). in western Poland, H. majus was observed in several ponds occurring within 2. Material and methods three active or abandoned fish farms and in a periodic pond at the bottom of the abandoned gravel pit. in total, Seeds for the study were obtained from herbarium the species was found in 5 localities in western Poland specimens representing populations of H. majus (nobis et al. 2018). distribu ted in western Poland (Table 1, Fig. 1). These Hypericum species occurring in Poland are mainly specimens are deposited in the herbarium of Depart- perennials, rarely bushes or annuals (among others, ment of Plant Taxonomy of Adam Mickiewicz Univer- Zarzycki et al. 2002; hypericum online). Their seeds are sity in Poznań (PoZ) (Fig. 2). of high diagnostic value in systematics and paleobotany Morphological structure was investigated based (robson 1981; hageman 1987; Meseguer & Sanmartín on the dry seed analysis using a stereomicroscope and 2012; Alonso et al. 2013). A particularly important scanning electron microscope (SeM). For SeM obser- feature is the sculpture of the seed coat and cell arrange- vations, 5 seeds were sampled and coated with gold. ment of its outer layer. however, only a relatively small All morphological measurements were made using number of Hypericum species have been studied so far a stereomicroscope with NIS-Elements Microscope in terms of seed morphology (Meseguer & Sanmartín Imaging Software. SeM images were used in analysis 2012; Alonso et al. 2013; Szkudlarz & Celka 2016) and of quantitative traits of the seed coat. in this work, we Fig. 1. Distribution of Hypericum majus in western Poland in the ATPol grid squares of 5×5 km biodiv. res. Conserv. 55: 7-14, 2019 9 Table 1. origin of the examined seed samples of Hypericum majus locality geographic habitat ATPol Data zbioru Collector coordinates rościce village, Żarskie 51°36`24`` n alt. 160 m, in 3 ponds AD8503 2 August 2017 S. rosadziński hills 14°59`38`` e on a fishing farm Miłowice village, Żarskie 51°36`5,37`` n alt. 147 m, in an AD8504 3 August 2017 S. rosadziński hills 15°3`18,26`` e overgrowing pond near bobrowice village, 51°31`17`` n alt. 137 m, in an AD9804 18 August 2017 S. rosadziński Dolnośląskie Forest 15°31`30`` e overgrowing pond near boruszyn village, 51°36`41,01`` n alt. 145 m, in an AD8503 16 november 2018 S. rosadziński Żarskie hills 14°58`23,50`` e overgrowing pond near Żary city, Żarskie 51°37`43,04`` n alt. 157 m, in an AD8503 16 november 2018 S. rosadziński hills 15°5`23,03`` e abandoned gravel pit Fig. 2. herbarium voucher of Hypericum majus from herbarium of the Department of Plant Taxonomy, Adam Mickiewicz University in Poznań (PoZ) 10 Piotr Szkudlarz et al. Seed morphology and anatomy of Hypericum majus (A. gray) britton followed terminology widely used in the studies on seed morphology (robson 1981; bojňanský & Fargašová 2007; Meseguer & Sanmartín 2012). To investigate the seed anatomical structure, ma- terial was subjected to a microtechnique procedure. Seeds were placed in the mixture of 2% solutions of glutaraldehyde and paraformaldehyde, then, they were kept for 1h in vacuum chamber (0.2 atm) and, next, in fridge (4oC) overnight. Fixed seeds were three times rinsed in 0.05 M cacodylate buffer and dehydrated in the series of ethanol solutions with increasing concentration (5, 10, 30, 50, 70, 80, 90, 96 and 100%). To prepare Fig. 4. Hypericum majus, seed (lM) microscopic slides, Technovit 7100 acrylic resin was used (heraeus Kulzer gmbh, Wahrheim, germany) as an embedding medium. Slices of 4 µm in thickness were stained with 0.05% solution of toluidine blue prepared in 1% sodium tetraborate. Additionally, seeds were stained with methylene blue-azure ii and basic fuchsine prepared for the staining of epoxy-embedded tissue sections (humphrey & Pitmann 1974). Slides were analysed under the AxioScopeA1 microscope. Lignin detection in the walls of seed coat cells was conducted under the AxioScopeA1 microscope with equipped Colibri 4. Unstained microscopic slides were observed using excitation light of 485 nm. For this wavelength, autofluorescence of lignin shows up as green light emission within visible spectrum. Fig. 5. Hypericum majus, seed (SeM), an abdominal view Photographic documentation of all slides was made using the AxioCam Mrc5 digital camera (Carl Zeiss gmbh, Jena, germany). there is a narrow, longitudinal scar after funiculus on 3. Results the incurved side (Fig. 4-5). Under a light microscope, their surface is lustrous. Seeds are light brown (honey 3.1. Morphological structure shade) in colour (Fig. 3). Under a SeM microscope, The studied seeds of H.
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